RESEARCH ARTICLE Molecular Genealogy of a Mongol Queen’s Family and Her Possible Kinship with Genghis Khan Gavaachimed Lkhagvasuren1, Heejin Shin2, Si Eun Lee2, Dashtseveg Tumen3, Jae- Hyun Kim4, Kyung-Yong Kim5, Kijeong Kim5, Ae Ja Park6, Ho Woon Lee2, Mi Jin Kim2, Jaesung Choi2, Jee-Hye Choi2, Na Young Min2, Kwang-Ho Lee1,2* 1 Department of Science of Cultural Heritage, Graduate School, Chung-Ang University, Seoul, Korea, 2 Department of Life Science, College of Natural Sciences, Chung-Ang University, Seoul, Korea, a11111 3 Department of Anthropology and Archaeology, School of Social Science, National University of Mongolia, Ulaanbaatar, Mongolia, 4 Department of Archaeology and Art History, College of Humanities, Donga University, Busan, Korea, 5 Institute for Medical Sciences, College of Medicine, Chung-Ang University, Seoul, Korea, 6 Department of Laboratory Medicine, Chung-Ang University College of Medicine, Seoul, Korea * [email protected] OPEN ACCESS Citation: Lkhagvasuren G, Shin H, Lee SE, Tumen Abstract D, Kim J-H, Kim K-Y, et al. (2016) Molecular Genealogy of a Mongol Queen’s Family and Her Members of the Mongol imperial family (designated the Golden family) are buried in a secret Possible Kinship with Genghis Khan. PLoS ONE necropolis; therefore, none of their burial grounds have been found. In 2004, we first discov- 11(9): e0161622. doi:10.1371/journal.pone.0161622 ered 5 graves belonging to the Golden family in Tavan Tolgoi, Eastern Mongolia. To define Editor: David Caramelli, University of Florence, the genealogy of the 5 bodies and the kinship among them, SNP and/or STR profiles of ITALY mitochondria, autosomes, and Y chromosomes were analyzed. Four of the 5 bodies were Received: November 4, 2015 determined to carry the mitochondrial DNA haplogroup D4, while the fifth carried hap- Accepted: July 12, 2016 logroup CZ, indicating that this individual had no kinship with the others. Meanwhile, Y-SNP and Y-STR profiles indicate that the males examined belonged to the R1b-M343 hap- Published: September 14, 2016 logroup. Thus, their East Asian D4 or CZ matrilineal and West Eurasian R1b-M343 patrilin- Copyright: © 2016 Lkhagvasuren et al. This is an eal origins reveal genealogical admixture between Caucasoid and Mongoloid ethnic open access article distributed under the terms of the Creative Commons Attribution License, which permits groups, despite a Mongoloid physical appearance. In addition, Y chromosomal and autoso- unrestricted use, distribution, and reproduction in any mal STR profiles revealed that the four D4-carrying bodies bore the relationship of either medium, provided the original author and source are mother and three sons or four full siblings with almost the same probability. Moreover, the credited. geographical distribution of R1b-M343-carrying modern-day individuals demonstrates that Data Availability Statement: All human remains descendants of Tavan Tolgoi bodies today live mainly in Western Eurasia, with a high fre- used in this manuscript are publicly available with the restricted use, because ancient human skeletons quency in the territories of the past Mongol khanates. Here, we propose that Genghis Khan excavated in Mongolia are regarded as cultural and his family carried Y-haplogroup R1b-M343, which is prevalent in West Eurasia, rather assets in Mongolia and are limited in the amount than the Y-haplogroup C3c-M48, which is prevalent in Asia and which is widely accepted to available for scientific research. Accordingly, be present in the family members of Genghis Khan. Additionally, Tavan Tolgoi bodies may researchers who are interested in using Tavan Tolgoi ’ samples need to obtain the approval of the Mongolian have been the product of marriages between the lineage of Genghis Khan s Borjigin clan government for foreign transfer of those samples, and and the lineage of either the Ongud or Hongirad clans, indicating that these individuals were to obtain the permission of authors, K-H Lee and D. members of Genghis Khan’s immediate family or his close relatives. Tumen. Mongolian Ministry of Education and Science address: Baga Toiruu-44, Government building 3, Sukhbaatar District, Ulaanbaatar City, Mongolia PLOS ONE | DOI:10.1371/journal.pone.0161622 September 14, 2016 1/31 Molecular Genealogy of a Mongol Queen's Family Phone: +976 1800-1209 Fax: +976 (11) 323158 Introduction E-mail: [email protected] Web site: http://www. meds.gov.mn/frontpage Mongolian National Temujin was born into the Borjigin clan as a son of Yesugei, who was a grandson of Khabul (or Inspection center address: Barilgachidiin Talbai-3, Qabul) Khan (King in the Mongolian language), the first khan of the Khamag Mongol confed- Government building 12, Chingeltei District, eration. In 1206, Temujin annexed and unified many Mongol-Turkic nomadic tribes of North- Ulaanbaatar City, Mongolia Phone: +976 (51) east Asia, and was then crowned the “Genghis Khan (the supreme king in the Mongolian 264786, +976 1800-1286 E-mail: info@inspection. language)” at a Kurultai, a general council of Mongol chiefs [1]. After founding the Mongol gov.mn Web site: www.inspection.gov.mn. Empire, Genghis Khan invaded neighboring lands outward from the Mongolian plateau, ulti- Funding: This work was supported by the National mately conquering most of Eurasia. The Mongol Empire expanded to be the largest contiguous Research Foundation of Korea (NRF) grant funded land empire in human history, covering the area from Eastern Europe to the East Sea/the Sea by the Korea government (MEST) (No. 2012R1A2A2A03047236) (www.nrf.re.kr/). This of Japan. The vast transcontinental empire allowed for the exchange of cultures and religions research was also supported by the Chung-Ang between Asia and Europe via the Silk Road. Thus, the Pax Mongolica greatly influenced many University Graduate Research Scholarship in 2015. civilizations in Eurasia during the 13th and 14th centuries; indeed, its cultural, social, religious, Competing Interests: The authors have declared and economic impact on the world remains today. that no competing interests exist. To solidify the foundation of the Mongol Empire, Genghis Khan employed two major strat- egies. First, because the Mongol Empire was too large to be controlled by a single ruler, he allo- cated the territories to his family members and let them rule their own independent territories [2, 3]. Because his wives were old and his sons were incompetent compared to his daughters, Genghis Khan bestowed upon his daughters, instead of his sons, the heavy responsibilities of shielding the inner territory of the Mongolian plateau and operating the outposts for his world conquest. He distributed the neighboring kingdoms surrounding the inner territory of the Mongolian plateau among his 4 daughters, including Alaqai Beki, “beki” meaning princess in the Mongolian language, who came to dominate the Ongud kingdom, Eastern Mongolia [2, 4]. His daughters faithfully ruled their own kingdoms throughout their lives on behalf of their father. Genghis Khan’s second strategy was to use quda, the traditional marriage alliance system of Mongolia, to marry his sons and daughters into the ruling lineages of neighboring kingdoms such as the Ongud [3, 5]. Through this system, Genghis Khan expected his daughters to become regents of the kingdoms once dominated by their husbands (guregens; prince consorts in the Mongolian language); he forced guregens to go to war, leaving their wives (bekis) in charge of running the home according to Mongol tradition. In doing so, Genghis Khan con- ferred power to his daughters, and not to the guregens, to rule the kingdoms. Moreover, gure- gens could not return to their homeland for long periods of time, and were killed at a high rate in Genghis Khan’s war. Through these qudas, bekis, as authoritarian rulers, strengthened alli- ances among their kingdoms and provided Genghis Khan the solid foundation necessary to conquer many kingdoms outward from the Mongol steppe [2]. Thus, Genghis Khan could not have founded the Mongol Empire without his bekis domi- nating the kingdoms [2, 3]. Although Genghis Khan’s daughters wielded unprecedented politi- cal authority in several kingdoms of the Mongol Empire, their names and achievements in solidifying the Mongol Empire have disappeared from Mongol chronicles over generations. Their burial grounds have never been found because all members of the imperial family of the Mongol Empire, including khans, khatuns, meaning “empress” in Mongolian, bekis, and their descendants, were buried without identifying signs, according to the long-standing tradition of keeping the burial grounds of ancestors in a secret necropolis called “Lord’s Enclosure” [3–5]. Indeed, the geographical locations of the graves of the Mongol imperial family (designated the Golden family) members are unknown [1, 6, 7]. Many researchers believe that the discovery of graves of the ancients will undoubtedly reveal details of their genealogies and lives. Molecular archaeologists have developed scientific and systematic approaches to trace customs, diseases, and genealogies of the ancients as well as PLOS ONE | DOI:10.1371/journal.pone.0161622 September 14, 2016 2/31 Molecular Genealogy of a Mongol Queen's Family various activities during the lifetimes of ancient peoples. Accordingly, it is likely that direct molecular archaeological analysis of the human remains from Genghis Khan’s Golden family members who ruled Pax Mongolica will provide scientific clues to unveil their mysterious lives and genealogies. In 2004, 7 graves were first excavated in the central hill of Tavan Tolgoi (“five hills” in the Mongolian language) by a Mongolian excavation team. Tavan Tolgoi lies within the Ongud province once dominated by Alaqai Beki and then Sorkhokhtani, a wife of Genghis Khan’s youngest son Tolui, during the early Mongolian era [8]. Mongolian archaeologists who partici- pated in the excavation of Tavan Tolgoi graves strongly suspected that 5 of 7 Tavan Tolgoi graves belonged to the Golden family [9–11], and one of those 5 graves was thought to be that of a Mongol Queen.